Lepus europaeus Pallas 1778

• Main Author: Bock, Anni
• Format: Text
• Language: unknown
• Published: Zenodo 2020
• Subjects: Biodiversity; Taxonomy; Animalia; Chordata; Mammalia; Lagomorpha; Leporidae; Lepus; Lepus europaeus; Arctic; Canada; Indian; New Zealand; Argentina; Wittenberg; Uruguay; Blanchard; Burrows; Myers; Morrison; Barrett; Bray; Genova; Browning; Verdier; Fang; Gomez; Jennings; Lubbock; Bertrand; Goodwin; Kos; Nowak; Woodstock; Meunier; Kowalczyk; Derevnya; Lepus timidus; mountain hare; Siberia
• Online Access: https://dx.doi.org/10.5281/zenodo.4570408; https://zenodo.org/record/4570408

Lepus europaeus Pallas, 1778 European Hare Lepus (Eulagos) europaeus europaeus Pallas, 1778:30. Type locality not stated; restricted to “southern Poland” by Trouessart (1910); further restricted to southwestern Poland by Ognev (1940:140). Lepus t [ imidus ]. albus Bechstein, 1801:1096. Type locality “Thuringia, Germany.” Lepus t [ imidus ]. flavus Bechstein, 1801:1096. Type locality “Thuringia, Germany.” Lepus t [ imidus ]. niger Bechstein, 1801:1096. Type locality “Thuringia, Germany.” Lepus medius Nilsson, 1820:224: Type locality “Zealand, Denmark.” Lepus hybridus Desmarest, 1822:349. Type locality “certain cantons of Siberia and Russia.” Lepus syriacus Hemprich and Ehrenberg, 1832: pt. 2, sig. u, fig, XIV, II. Type locality “Mount Lebanon.” Lepus caspicus Hemprich and Ehrenberg, 1832: pt. 2, sig. y. Type locality “Astrachan [=Asktrakhan, Russia].” Lepus aquilonius Blasius, 1842:89. Type locality “ hauptsächlich den Gegenden zwischen dem 56.° und 63. ° N. Br.;” central Russia. Lepus timidus var. hymenalis Tumac Eversmann, 1850:201. Type locality “Kazan.” Lepus campicola Gervais, 1859:47. Nomen nudum. Lepus meridionalis Gervais, 1859:47. Nomen nudum. Eulagos Judeae Gray, 1867:222. Type locality “The Holy Land;” restricted to Ain Faskhah, Palestine by Qumsiyeh (1996:314). Lepus timidus coronatus Fitzinger, 1867:161. Nomen nudum. Lepus timidus rufus Fitzinger, 1867:161. Nomen nudum. Lepus timidus cinereus Fitzinger, 1867:161. Nomen nudum. Lepus timidus nigricans Fitzinger, 1867:161. Nomen nudum. Lepus timidus maculatus Fitzinger, 1867:161. Nomen nudum. Lepus campestris Bogdanov, 1871:175. Name preoccupied, not L . campestris Bachman, 1837. Lepus timidus var. argenteogrisea König-Warthausen, 1875:277. Type locality “Ulm,” Württemberg, Germany. Lepus timidus var. tumak Tichomirov and Korchagin, 1889:31. First suitable name for tumac Eversmann, 1850. Lepus transsylvanicus Matschie, 1901:236. Type locality “von Taslau in Rumänien.” Lepus creticus Barrett-Hamilton, 1903:126. Type locality “Crete.” Lepus cyprius Barrett-Hamilton, 1903:127. Type locality “Cyprus.” Lepus parnassius Miller, 1903:145. Type locality “Agorianni, north side of Lyakura (Parnassus) Mts., Greece.” Lepus cyrensis Satunin, 1905:60. Type locality “Village Barda, Djevanshirski county, Elisavetpolski province of Russia.” Lepus e [ uropaeus ]. pyrenaicus Hilzheimer, 1906:512. Type locality “(Bagnèrs), Pyrenäen.” © The Author(s) 2020. Published by Oxford University Press on behalf of the American Society of Mammalogists, www.mammalogy.org. 125 Lepus e [ uropaeus ]. transsylvaticus Hilzheimer, 1906:512. Incorrect subsequent spelling of transsylvanicus Matschie, 1901. Lepus e [ uropaeus ]. meridiei Hilzheimer, 1906:512. Type locality “(Aveyron), Sudfrankreich [=southern France].” Lepus dayanus connori Robinson, 1918:15. Type locality “between Ahwaz and Mohammerah, Karun R. [=Karun River], Persia.” Lepus europaeus tesquorum Ognev (in Ognev and Worobiev, 1923), 1923:115. Type locality “Dokuchaevskaya Experimental Station.” Lepus europaeus ghigi de Beaux, 1927:2. Type locality “Stampalia, Island, Aegean Sea” vide Ellerman and Morrison-Scott (1966:437). Lepus europaeus caucasicus Ognev, 1929.:75. Type locality “Umgebung von Wladikawkas [=vicinity of Vladikavkaz, North Ossetia].” L [ epus ]. e [ uropaeus ]. caucasicus nato ponticus Ognev, 1929:75. Type locality “Ansiedlung Beta, Tschernomorski-Gouvernement,” Black Sea Coast. L [ epus ]. e [ uropaeus ]. caspicus nato kalmykorum Ognev, 1929:75. Type locality “Sangegend [= sand zone of] Akima Ikizochura, Kalmyken-Steppe.” Lepus europaeus carpathous de Beaux, 1929:144. Type locality “a Scarpanto [=in Karpathos]. Lepus europaeus iranensis Goodwin, 1939:4. Type locality “ Teheran, Iran; alt about 3500 feet.” Lepus europaeus niethammeri Wettstein, 1943:282. Type locality “Vytina, 1000m altitude, Peloponnese,” southern Greece. Lepus europaeus borealis Kuznetsov, 1944:271. Type locality “Northern Bashkiria;” Bashkortostan Republic of Russia. Lepus europaeus biarmicus Heptner, 1948:709. Replacement name for Lepus europaeus borealis Kuznetsov, 1944. Lepus cyanotus Blanchard, 1957:30. Type locality “Lanslebourg, Savoie, France, 2600 m.” Lepus capensis astaricus Baloutch, 1978:444. Type locality “dans la forêt d’Astara à 30 km de la Mer Caspienne.” CONTEXT AND CONTENT. Order Lagomorpha, family Leporidae, genus Lepus . Sixteen subspecies are currently recognized (Schai-Braun and Hackländer 2016): L. e. caspicus Hemprich and Ehrenberg, 1832. See above; kalmykorum Ognev is a synonym. L. e. connori Robinson, 1918. See above; astaricus Baloutch and iranensis Goodwin are synonyms. L. e. creticus Barrett-Hamilton, 1903. See above. L. e. cyprius Barrett-Hamilton, 1903. See above. L. e. cyrensis Satunin, 1905. See above; caucasicus Ognev, ghigi de Beaux, are synonyms. L. e. europaeus Pallas, 1778. See above; albus Bechstein, aregenteogrisea Konig-Warthausen, cyanotus Blachard, flavus Bechstein, niger Bechstein, and pyrenaicus Hilzheimer are synonyms. L. e. hybridus Desmarest, 1822. See above; aquilonius Blasius, biarmicus Heptner, borealis Kuznetsov, campestris Bogdanov, hyemalis Eversmann, tesquorum Ognev and Worobiev, tumac Tichomirov and Kortchagin are synonyms. L. e. judeae (Gray, 1867). See above. L. e. karpathorum Hilzheimer, 1906:512. Type locality “Karpathen [=Carpathian Mountains].” L. e. medius Nilsson, 1820. See above. L. e. occidentalis de Winton, 1898:152. Type locality “Herefordshire;” England. L. e. parnassius Miller, 1903. See above; niethammeri Wettstein is a synonym. L. e. ponticus Ognev, 1929. See above. L. e. rhodius Festa, 1914:9. Type locality “Island of Rhodes.” L. e. syriacus Hemprich and Ehrenberg, 1832. See above. L. e. transsylvanicus Matschie, 1901. See above; campicola Gervais, cinereus Fitzinger, coronatus Fitzinger, maculates Fitzinger, meridiei Hilzheimer, meridionalis Gervais, nigricans Fitzinger, rufus Fitzinger, and transsylvaticus Hilzheimer are synonyms. NOMENCLATURE NOTES. The vernacular names of Lepus europaeus are European brown hare, European hare, brown hare, and field hare. In Germany field hare (Feldhase) is most commonly used throughout the country. Petter (1961) placed L. europaeus into L. capensis as a polytypic species which ranges from North East Africa eastward across the North Arabian Peninsula and the Middle East, and northward through Israel to Turkey, Russia, and China. He argued that body size, dental characters, and pelage color depend on climatic and ecological variation among habitats. This was accepted until further morphological studies argued the converse. Sludski and Strautmann (1980) stated the “large L. europaeus ” and “small L. capensis ” (= L. tolai ) in Kazakhstan were sympatric species without evidence of hybridization and interpreted it as overlapping ends of a Rassenkreis (ring of populations). Also results of reanalysis by Angermann (1983) indicated a marked discontinuity between both species running from the eastern Mediterranean coast (central Israel) through Iran, and on this basis separated L. europaeus from L. capensis and L. tolai (Hoffmann and Smith 2005). Recent research (Slimen et al. 2005, 2006, 2008) support Petter’s hypothesis that L. europaeus and L. capensis are conspecific. However, until conclusive evidence is available, most taxonomists retain L. europaeus and L. capensis as valid species (Hackländer and Schai-Braun 2018). DIAGNOSIS Lepus europaeus (Fig. 1) and the sympatric Lepus timidus (mountain hare) in the northern hemisphere are very similar in size.Although some arctic subspecies of L.timidus may be as large as, or even larger, than the largest L. europaeus , L. europaeus is generally heavier at 3.5–5.0 kg ( L . timidus averages from 2.4 kg in the Alps to 3.4 kg in Siberia—Angerbjorn 2018). Headbody length of L . europaeus is 550–650 mm (460–560 mm in L. timidus ). The ears are longer from notch 110–140 mm and when bent forward they are longer than forehead (versus 67–106 mm in L. timidus , ears when bent forward are equal in length to forehead); the tail is also longer at 75–140 mm (mean 95 mm) compared with 53–70 mm in L. timidus length of hind foot is 130–164 mm (mean 147 mm) which is shorter than in L . timidus (145–172 mm—Krapp 2003; Angerbjorn 2018). The length of the skull (from the most rostral to the most caudal point of the occipital) averages 97.1 mm in L . europaeus ( L. timidus averages 90 mm). The facial region of the skull is extended with markedly raised nasalia (Fig. 2) compared to a shorter facial and a more concave frontal region in L. timidus . Dentale is softer, processus condyloideus is inclined more posteriorly than in L. timidus . Diastema relatively long (average 31.2 mm), and at over 80% of lower toothrow, longer than in L. timidus , corpus mandibulae with incisivus (I) inclined anteroventrally (anterodorsal in L. timidus ). Incisivus 1 (I1) of L. europaeus are rectangular in section; ratio of length to width 0.74:0.80 compared to the more quadratic dimension in L. timidus of 0.84:0.95; incisors are additionally more curved than in L. timidus . Root of I1 far anterior of sutura premaxillo-maxillaris with a frequency of 78.3–100% compared to 98.6–100% in L. timidus . Zygomatic length (34.7–41.8 mm) and zygomatic width (44–49 mm) are longer than in L. timidus (length = 33.4–37.9; width = 42.5– 45.6 mm). Front and rear wings of supraorbital appendix are well developed and mostly free (not as well developed in L . timidus ), osseous palate shorter (compared to L . timidus ), choanae wide. Coat color of L. europaeus does not change to white in winter as in L. timidus , although animals in the northern latitudes may turn slightly lighter. Teat formula 2p + 4a = 6 compared to 2p + 6a = 8 for L. timidus (Krapp 2003). Although similar in appearance to L . europaeus , Lepus capensis (Cape hare) is overall smaller and lighter (with ear length 97–114 mm; skull length 27.5–30 mm; length of hind foot 93–103 mm, weight 1.5–2.5 kg). When compared to the sympatric Oryctolagus cuniculus (European rabbit) the difference is obvious. The European rabbit is smaller (body weight 0.9–1.8 kg, head–body length 375 [female] and 406 [male] mm), and has a more compact shape. The European rabbit burrows and its leverets are born nidicolous (altricial) compared to the L. europaeus which lives aboveground with nidifugous (precocial) leverets. Ears are shorter (when bent forward they do not extend to the nose) and have no black tips. Furthermore, the eyes do not have a light-brown iris but look black and the pelage has more grayish than yellowish base hair and longer guard hair with black ends (Zörner 1981; Krapp 2003). GENERAL CHARACTERS Lepus europaeus is the largest species of Lepus in Europe with a head–body length of 550–650 mm. The weight ranges from 3.5 to 5 kg and varies in both males and females during the course of the year, due to food supply and reproductive cycle. Length of cranium is 96–104 mm, width of zygomatic arch 44–51 mm, width of cranium 28.9–35.1 mm, cranium height 26.1–29.7 mm, length of lower toothrow 19–21 mm, and length of upper toothrow 17–19 mm. Depending on geographical location and seasonal changes, the pelage color of L. europaeus ranges from tawny-brown to grayish-brown, with a beige-white venter. The face is brown with beige eye rings. Eyes are large with black pupils and light-brown iris. Molting occurs twice a year, one starting around February the other starting in July. The winter coat may be slightly lighter toward yellowish-gray, especially in northern regions of its distribution. The tail (75–140 mm) is V-shaped, black above and white below. Length of hind foot is 130–150 mm. Ears with black tips are 110–140 mm, longer than the forehead when bent forward. External sexual dimorphism is lacking; thus, sex determination is only possible by examining genitalia (Zörner 1981; Krapp 2003). DISTRIBUTION During the last IceAge (about12,000 years ago) Lepus europaeus was forced into southern Europe, only L. timidus could survive in more northern habitats due to its adaptions to colder conditions. After that L. europaeus recolonized the postglacial steppe landscape and later adapted well to agricultural farmland (Fickel 2003). Lepus europaeus is native in Albania, Austria, Belarus, Belgium, Bosnia and Herzegovina, Bulgaria, Croatia, Czech Republic, Denmark, Estonia, Finland, France, Germany, Greece, Hungary, Islamic Republic of Iran, Iraq, Israel, Italy, Latvia, Liechtenstein, Lithuania, Luxembourg, Macedonia, Moldavia, Montenegro, Netherlands, Poland, Romania, Russian Federation, Serbia, Slovakia, Slovenia, Spain, Switzerland, Syrian Arab Republic, Turkey, and Ukraine. L. europaeus was introduced in Argentina, Australia and Tasmania, Barbados, Brazil, Canada, Chile, Crete, Falkland Islands (Malvinas), Ireland, New Zealand, Réunion, Sweden, United Kingdom, United States, and Uruguay (Fig. 3; Hackländer and Schai-Braun 2019). Masseti (2012) gives a broad overview of several Lepus taxa of the Ionian and Aegean Islands. He lists L. e. creticus , from Cephalonia, Crete, and several small islets of Crete, including Gavdhos, Gavdhopoula, Gaidouronisi (Chrysi), Koufonisi, and the Archipelago of the Dionysades. Furthermore, L. e. ghigii , from Astypalaia, L. e. carpathous , from Karpathos, L. e. rhodius , from Rhodos, and L. e. cyrensis , from the island of Kos. Subspecies of L. europaeus of Paximada (Dionysades Islands) and Koufonisi are smaller than those of Crete; however, they fit the typical morphology of Lepus europaeus europaeus , and not L. capensis (Niethammer 1992). Distinction of insular subspecies was based on coat coloration, body, and skull measurements, and they can be regarded more as ecophenotypic varieties than true geographical subspecies. Nonetheless, Mamuris et al. (2001) found high mtDNA variability and detected haplotypes from the Greek hares that were different from those of Central Europe. Based on this, Suchentrunk et al. (2003) analyzed the multilocus allozyme variability of Greek hares with regard to the occurrence of indigenous alleles due to the late Pleistocene refuge (see “Genetics”). In summary, Greek hares differ from Bulgarian hares, but within Greek hares the gene pool diversity is slight, thus favoring the hypothesis of a late Pleistocene refugial population in the southern Balkans, with little genetic differentiation among Greek subspecies to support earlier descriptions. A clear difference can be made between Anatolian and European brown hares. In Turkey, Demirbaş and Albayrak (2014) compared morphometric data and pelage color of L. e. europaeus subspecies, namely, L. e. transsylvanicus , L. e. cyrensis , L. e. caucasicus , and L. e. syriacus , and concluded that Anatolian hares belong to the subspecies L. e. syriacu s. In Canada, L. e. hybridus was introduced and established in Ontario around the Great Lakes and south of the Canadian Shield. Nine individuals escaped from captivity near Brantford, Ontario, in 1912, slowly spreading in southern Ontario north of Lake Erie (St. Thomas and Woodstock), west and north of Lake Ontario (Toronto) to Goodrich on the east side of Lake Huron. However, it has failed to spread further north (Reynolds and Stinson 1959; Whitaker and Hamilton 1998). In the United States L. e. hybridus was introduced into Dutchess County, New York, in 1893. Importations continued through 1911 to provide a game species. L. e. hybridus was introduced and established in New York and Connecticut (Hall 1951; Whitaker and Hamilton 1998). In Australia L. europaeus was introduced and became established on the shores of Westernport Bay, Victoria in 1862. Another colony was set up in 1863 on Phillip Island by the Acclimatization Society of Victoria where it thrived and became a source for distribution to many parts, for example, Tasmania and New Zealand (Myers et al. 1989). In Australia L. europaeus now occupies about 700,000 km 2 in the east and south (Stott 2003). In New Zealand the first L. europaeus introduction was in 1851 from Britain but it died out, the second introduction of six L. europaeus came also from Britain via Australia in 1863 and established a stable population (up to 200 in 1865). Compared to European rabbits introduced at around the same time it did not become a pest (Flux 1990). For South America, Grigera and Rapoport (1983) and Bonino et al. (2010) give detailed but somewhat conflicting information about L. europaeus (considering it conspecific with L. capensis ). Thirty-six L. europaeus were imported from Germany in 1888 at “Estancia La Hansa” near the town of Cañada de Gomez, Province of Santa Fe, Argentina. From France L. europaeus was introduced in 1897 near Tandil, Province of Buenos Aires, Argentina, and nine L. europaeus were liberated in the Province of Santa Cruz, southern Argentina in 1930. In the 1980s, L. europaeus occupied large parts of Argentina, Chile, Uruguay, Paraguay, and southern parts of Bolivia and Brazil. Since then it spread further north and, probably coming from Bo : Published as part of Bock, Anni, 2020, Lepus europaeus (Lagomorpha: Leporidae), pp. 125-142 in Mammalian Species 52 (997) on pages 125-138, DOI: 10.1093/mspecies/seaa010, http://zenodo.org/record/4570375 : {"references": ["PALLAS, P. S. 1778. Novae Species Quadrupedum e Glirium Ordine. SVMTV Wolfgang Waltheri, Erlangae, Germany.", "TROUESSART, E. L. 1910. Faune des Mammifres D'Europe. R. Friedlander & Sohn, Berlin, Germany.", "OGNEV, S. I. 1940. Zveri SSSR i prilezhashchikh stran: Gryzuny. (Zveri vostochnoiEvropy i severnoi Azii) [Mammals of the USSR and adjacent countries: rodents (mammals of eastern Europe and northern Asia)]. Akademiya Nauk SSSR 4: 1 - 615 (in Russian).", "BECHSTEIN, J. M. 1801. 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